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Mollusca
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which
Mollusca
are known as molluscs or mollusks[a] (/ˈmɒləsk/). Around 85,000  extant species of molluscs are
recognized.[3] The number of fossil species is estimated between 60,000 and 100,000 additional species.[4] Temporal range:
The proportion of undescribed species is very high. Many taxa remain poorly studied.[5]

Molluscs are the largest marine phylum, comprising about 23% of all the named marine organisms.
Numerous molluscs also live in freshwater and terrestrial habitats. They are highly diverse, not just in size
and anatomical structure, but also in behaviour and habitat. The phylum is typically divided into 7 or
8[6] taxonomic classes, of which two are entirely extinct. Cephalopod molluscs, such as squid, cuttlefish, and
octopuses, are among the most neurologically advanced of all invertebrates—and either the giant squid or
the colossal squid is the largest known invertebrate species. The gastropods (snails and slugs) are by far the
most numerous molluscs and account for 80% of the total classified species. Clockwise from top-left: Helix pomatia, a
gastropod; Common octopus, a
The three most universal features defining modern molluscs are a mantle with a significant cavity used for
breathing and excretion, the presence of a radula (except for bivalves), and the structure of the nervous cephalopod; Tonicella lineata, a chiton;
system. Other than these common elements, molluscs express great morphological diversity, so many Atlantic surf clam, a bivalve.
textbooks base their descriptions on a "hypothetical ancestral mollusc" (see image below). This has a single,
Scientific classification
"limpet-like" shell on top, which is made of proteins and chitin reinforced with calcium carbonate, and is
secreted by a mantle covering the whole upper surface. The underside of the animal consists of a single Kingdom: Animalia
muscular "foot". Although molluscs are coelomates, the coelom tends to be small. The main body cavity is a
Subkingdom: Eumetazoa
hemocoel through which blood circulates; as such, their circulatory systems are mainly open. The
"generalized" mollusc's feeding system consists of a rasping "tongue", the radula, and a complex digestive Clade: ParaHoxozoa
system in which exuded mucus and microscopic, muscle-powered "hairs" called cilia play various important
Clade: Bilateria
roles. The generalized mollusc has two paired nerve cords, or three in bivalves. The brain, in species that
have one, encircles the esophagus. Most molluscs have eyes, and all have sensors to detect chemicals, Clade: Nephrozoa
vibrations, and touch. The simplest type of molluscan reproductive system relies on external fertilization,
but more complex variations occur. Nearly all produce eggs, from which may emerge trochophore larvae, (unranked): Protostomia
more complex veliger larvae, or miniature adults. The coelomic cavity is reduced. They have an open (unranked): Spiralia
circulatory system and kidney-like organs for excretion.
Superphylum: Lophotrochozoa
Good evidence exists for the appearance of gastropods, cephalopods, and bivalves in the Cambrian period, Phylum: Mollusca
541–485.4  million years ago. However, the evolutionary history both of molluscs' emergence from the
Linnaeus, 1758
ancestral Lophotrochozoa and of their diversification into the well-known living and fossil forms are still
subjects of vigorous debate among scientists. Classes

Molluscs have been and still are an important food source for anatomically modern humans. Toxins that can
See text.
accumulate in certain molluscs under specific conditions create a risk of food poisoning, and many
jurisdictions have regulations to reduce this risk. Molluscs have, for centuries, also been the source of
important luxury goods, notably pearls, mother of pearl, Tyrian purple dye, and sea silk. Their shells have Diversity[1]
also been used as money in some preindustrial societies. 85,000 recognized living species.
A handful of mollusc species are sometimes considered hazards or pests for human activities. The bite of the
blue-ringed octopus is often fatal, and that of Octopus apollyon causes inflammation that can last over a month.
Stings from a few species of large tropical cone shells can also kill, but their sophisticated, though easily produced,
venoms have become important tools in neurological research. Schistosomiasis (also known as bilharzia,
bilharziosis, or snail fever) is transmitted to humans by water snail hosts, and affects about 200 million people.
Snails and slugs can also be serious agricultural pests, and accidental or deliberate introduction of some snail
species into new environments has seriously damaged some ecosystems.

Etymology
The words mollusc and mollusk are both derived from the French mollusque, which originated from the Latin Cornu aspersum (formerly Helix
molluscus, from mollis, soft. Molluscus was itself an adaptation of Aristotle's τὰ μαλάκια ta malákia (the soft aspersa) – a common land snail
ones; < μαλακός malakós "soft"), which he applied inter alia to cuttlefish.[7][8] The scientific study of molluscs is
accordingly called malacology.[9]

The name Molluscoida was formerly used to denote a division of the animal kingdom containing the
brachiopods, bryozoans, and tunicates, the members of the three groups having been supposed to somewhat
resemble the molluscs. As now known, these groups have no relation to molluscs, and very little to one another, so
the name Molluscoida has been abandoned.[10]

Definition
The most universal features of the body structure of molluscs are a mantle with a significant cavity used for
breathing and excretion, and the organization of the nervous system. Many have a calcareous shell.[11]

Molluscs have developed such a varied range of body structures, finding synapomorphies (defining
characteristics) to apply to all modern groups is difficult.[12] The most general characteristic of molluscs is they
are unsegmented and bilaterally symmetrical.[13] The following are present in all modern molluscs:[14][16] Fossilized ammonite displayed at
the National Museum of the
The dorsal part of the body wall is a mantle (or pallium) which secretes calcareous spicules, plates or shells. It Philippines
overlaps the body with enough spare room to form a mantle cavity.
The anus and genitals open into the mantle cavity.
There are two pairs of main nerve cords.[15]

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Other characteristics that commonly appear in textbooks have significant exceptions:

  Whether characteristic is found in these classes of Molluscs

Supposed
universal
Molluscan Aplacophora[15]: 291–292  Polyplacophora[15]: 292–298  Monoplacophora[15]: 298–300  Gastropoda[15]: 300–343  Cephalopoda[15]: 343–367  Bivalvia[15]: 36
characteristic[14]
Radula, a
rasping Absent in 20% of
Yes Yes Yes Yes No
"tongue" with Neomeniomorpha
chitinous teeth

Broad,
Reduced or absent Yes Yes Yes Modified into arms Yes
muscular foot

Dorsal
concentration of
Not obvious Yes Yes Yes Yes Yes
internal organs
(visceral mass)
Large digestive No ceca in some
Yes Yes Yes Yes Yes
ceca Aplacophora

Large complex
metanephridia None Yes Yes Yes Yes Yes
("kidneys")

Snails, yes; slugs, Octopuses, no;

One or more Primitive forms, yes;
Yes Yes mostly yes (internal cuttlefish, nautilus, Yes
valves/ shells modern forms, no
vestigial) squid, yes
Odontophore Yes Yes Yes Yes Yes No

Diversity
Estimates of accepted described living species of molluscs vary from 50,000 to a maximum of 120,000 species.[1]
The total number of described species is difficult to estimate because of unresolved synonymy. In 1969 David
Nicol estimated the probable total number of living mollusc species at 107,000 of which were about 12,000 fresh-
water gastropods and 35,000 terrestrial. The Bivalvia would comprise about 14% of the total and the other five
classes less than 2% of the living molluscs.[18] In 2009, Chapman estimated the number of described living
mollusc species at 85,000.[1] Haszprunar in 2001 estimated about 93,000 named species,[19] which include 23%
of all named marine organisms.[20] Molluscs are second only to arthropods in numbers of living animal species[17] Diversity and variability of shells of
molluscs on display
— far behind the arthropods' 1,113,000 but well ahead of chordates' 52,000.[15]: Front endpaper  About
200,000  living species in total are estimated,[1][21] and 70,000 fossil species,[14] although the total number of
mollusc species ever to have existed, whether or not preserved, must be many times greater than the number alive
today.[22]

Molluscs have more varied forms than any other animal phylum. They include snails, slugs and other gastropods;
clams and other bivalves; squids and other cephalopods; and other lesser-known but similarly distinctive
subgroups. The majority of species still live in the oceans, from the seashores to the abyssal zone, but some form a
significant part of the freshwater fauna and the terrestrial ecosystems. Molluscs are extremely diverse in tropical
and temperate regions, but can be found at all latitudes.[12] About 80% of all known mollusc species are
gastropods.[17] Cephalopoda such as squid, cuttlefish, and octopuses are among the neurologically most advanced About 80% of all known mollusc
of all invertebrates.[23] The giant squid, which until recently had not been observed alive in its adult form,[24] is species are gastropods (snails and
one of the largest invertebrates, but a recently caught specimen of the colossal squid, 10  m (33  ft) long and slugs), including this cowry (a sea
weighing 500 kg (1,100 lb), may have overtaken it.[25] snail).[17]

Freshwater and terrestrial molluscs appear exceptionally vulnerable to extinction. Estimates of the numbers of
nonmarine molluscs vary widely, partly because many regions have not been thoroughly surveyed. There is also a shortage of specialists who can
identify all the animals in any one area to species. However, in 2004 the IUCN Red List of Threatened Species included nearly 2,000 endangered
nonmarine molluscs. For comparison, the great majority of mollusc species are marine, but only 41 of these appeared on the 2004 Red List. About
42% of recorded extinctions since the year 1500 are of molluscs, consisting almost entirely of nonmarine species.[26]

Hypothetical ancestral mollusc

Anatomical diagram of a hypothetical ancestral mollusc

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Because of the great range of anatomical diversity among molluscs, many textbooks start the subject of molluscan anatomy by describing what is
called an archi-mollusc, hypothetical generalized mollusc, or hypothetical ancestral mollusc (HAM) to illustrate the most common features found
within the phylum. The depiction is visually rather similar to modern monoplacophorans.[12][16][27]

The generalized mollusc is an unsegmented, bilaterally symmetrical animal and has a single, "limpet-like" shell on top. The shell is secreted by a
mantle covering the upper surface. The underside consists of a single muscular "foot".[16] The visceral mass, or visceropallium, is the soft,
nonmuscular metabolic region of the mollusc. It contains the body organs.[13]

Mantle and mantle cavity

The mantle cavity, a fold in the mantle, encloses a significant amount of space. It is lined with epidermis, and is exposed, according to habitat, to sea,
fresh water or air. The cavity was at the rear in the earliest molluscs, but its position now varies from group to group. The anus, a pair of osphradia
(chemical sensors) in the incoming "lane", the hindmost pair of gills and the exit openings of the nephridia ("kidneys") and gonads (reproductive
organs) are in the mantle cavity.[16] The whole soft body of bivalves lies within an enlarged mantle cavity.[13]

Shell

The mantle edge secretes a shell (secondarily absent in a number of taxonomic groups, such as the nudibranchs[13]) that consists of mainly chitin and
conchiolin (a protein hardened with calcium carbonate),[16][28] except the outermost layer, which in almost all cases is all conchiolin (see
periostracum).[16] Molluscs never use phosphate to construct their hard parts,[29] with the questionable exception of Cobcrephora.[30] While most
mollusc shells are composed mainly of aragonite, those gastropods that lay eggs with a hard shell use calcite (sometimes with traces of aragonite) to
construct the eggshells.[31]

The shell consists of three layers: the outer layer (the periostracum) made of organic matter, a middle layer made of columnar calcite, and an inner
layer consisting of laminated calcite, often nacreous.[13]

In some forms the shell contains openings. In abalones there are holes in the shell used for respiration and the release of egg and sperm, in the
nautilus a string of tissue called the siphuncle goes through all the chambers, and the eight plates that make up the shell of chitons are penetrated
with living tissue with nerves and sensory structures.[32]

Foot

The underside consists of a muscular foot, which has adapted to different purposes (locomotion, grasping the
substratum, burrowing or feeding) in different classes.[33] The foot carries a pair of statocysts, which act as
balance sensors. In gastropods, it secretes mucus as a lubricant to aid movement. In forms having only a top shell,
such as limpets, the foot acts as a sucker attaching the animal to a hard surface, and the vertical muscles clamp
the shell down over it; in other molluscs, the vertical muscles pull the foot and other exposed soft parts into the
shell.[16] In bivalves, the foot is adapted for burrowing into the sediment;[33] in cephalopods it is used for jet
propulsion,[33] and the tentacles and arms are derived from the foot.[34] 0:51

A 50-second video of snails (most

Circulatory system
likely Natica chemnitzi and
Cerithium stercusmuscaram)
Most molluscs' circulatory systems are mainly open, except for cephalopods, whose circulatory systems are closed.
feeding on the sea floor in the Gulf
Although molluscs are coelomates, their coeloms are reduced to fairly small spaces enclosing the heart and of California, Puerto Peñasco,
gonads. The main body cavity is a hemocoel through which blood and coelomic fluid circulate and which encloses Mexico
most of the other internal organs. These hemocoelic spaces act as an efficient hydrostatic skeleton.[13] The blood
of these molluscs contains the respiratory pigment hemocyanin as an oxygen-carrier. The heart consists of one or
more pairs of atria (auricles), which receive oxygenated blood from the gills and pump it to the ventricle, which pumps it into the aorta (main artery),
which is fairly short and opens into the hemocoel.[16] The atria of the heart also function as part of the excretory system by filtering waste products
out of the blood and dumping it into the coelom as urine. A pair of nephridia ("little kidneys") to the rear of and connected to the coelom extracts any
re-usable materials from the urine and dumps additional waste products into it, and then ejects it via tubes that discharge into the mantle cavity.[16]

Exceptions to the above are the molluscs Planorbidae or ram's horn snails, which are air-breathing snails that use iron-based hemoglobin instead of
the copper-based hemocyanin to carry oxygen through their blood.

Respiration

Most molluscs have only one pair of gills, or even only a singular gill. Generally, the gills are rather like feathers in shape, although some species have
gills with filaments on only one side. They divide the mantle cavity so water enters near the bottom and exits near the top. Their filaments have three
kinds of cilia, one of which drives the water current through the mantle cavity, while the other two help to keep the gills clean. If the osphradia detect
noxious chemicals or possibly sediment entering the mantle cavity, the gills' cilia may stop beating until the unwelcome intrusions have ceased. Each
gill has an incoming blood vessel connected to the hemocoel and an outgoing one to the heart.[16]

Eating, digestion, and excretion

Molluscs use intracellular digestion. Most molluscs have muscular mouths with radulae, "tongues", bearing many rows of chitinous teeth, which are
replaced from the rear as they wear out. The radula primarily functions to scrape bacteria and algae off rocks, and is associated with the odontophore,
a cartilaginous supporting organ.[13] The radula is unique to the molluscs and has no equivalent in any other animal.

Molluscs' mouths also contain glands that secrete slimy mucus, to which the food sticks. Beating cilia (tiny "hairs") drive the mucus towards the
stomach, so the mucus forms a long string called a "food string".[16]

At the tapered rear end of the stomach and projecting slightly into the hindgut is the prostyle, a backward-pointing cone of feces and mucus, which is
rotated by further cilia so it acts as a bobbin, winding the mucus string onto itself. Before the mucus string reaches the prostyle, the acidity of the
stomach makes the mucus less sticky and frees particles from it.[16]

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The particles are sorted by yet another group of cilia, which send the smaller particles, mainly minerals, to the
prostyle so eventually they are excreted, while the larger ones, mainly food, are sent to the stomach's cecum (a
pouch with no other exit) to be digested. The sorting process is by no means perfect.[16]

Periodically, circular muscles at the hindgut's entrance pinch off and excrete a piece of the prostyle, preventing
the prostyle from growing too large. The anus, in the part of the mantle cavity, is swept by the outgoing "lane" of
the current created by the gills. Carnivorous molluscs usually have simpler digestive systems.[16]
Snail radula at work
As the head has largely disappeared in bivalves, the mouth has been equipped with labial palps (two on each side    = Food        = Radula
of the mouth) to collect the detritus from its mucus.[13]    = Muscles
   = Odontophore "belt"

Nervous system

The cephalic molluscs have two pairs of main nerve cords organized around a number of paired ganglia, the
visceral cords serving the internal organs and the pedal ones serving the foot. Most pairs of corresponding ganglia
on both sides of the body are linked by commissures (relatively large bundles of nerves). The ganglia above the gut
are the cerebral, the pleural, and the visceral, which are located above the esophagus (gullet). The pedal ganglia,
which control the foot, are below the esophagus and their commissure and connectives to the cerebral and pleural
ganglia surround the esophagus in a circumesophageal nerve ring or nerve collar.[16]

The acephalic molluscs (i.e., bivalves) also have this
have only three pairs of ganglia— cerebral, pedal,
important of the three functioning as the principal
around the edges of their shells which connect to
distinguish between light and shadow.
ring but it is less obvious and less important. The bivalves
and visceral— with the visceral as the largest and most
center of "thinking". Some such as the scallops have eyes
a pair of looped nerves and which provide the ability to

Simplified diagram of the mollusc Reproduction

nervous system
The simplest molluscan reproductive system relies on external Apical tuft (cilia)
fertilization, but with more complex variations. All produce eggs, from
which may emerge trochophore larvae, more complex veliger larvae, or miniature adults. Two gonads sit Prototroch (cilia)
next to the coelom, a small cavity that surrounds the heart, into which they shed ova or sperm. The Stomach
nephridia extract the gametes from the coelom and emit them into the mantle cavity. Molluscs that use such Mouth
a system remain of one sex all their lives and rely on external fertilization. Some molluscs use internal Metatroch (cilia)
fertilization and/or are hermaphrodites, functioning as both sexes; both of these methods require more Mesoderm
complex reproductive systems.[16] Anus
/// = cilia
The most basic molluscan larva is a trochophore, which is planktonic and feeds on floating food particles by Trochophore larva[35]
using the two bands of cilia around its "equator" to sweep food into the mouth, which uses more cilia to
drive them into the stomach, which uses further cilia to expel undigested remains through the anus. New
tissue grows in the bands of mesoderm in the interior, so the apical tuft and anus are pushed further apart as the animal grows. The trochophore
stage is often succeeded by a veliger stage in which the prototroch, the "equatorial" band of cilia nearest the apical tuft, develops into the velum
("veil"), a pair of cilia-bearing lobes with which the larva swims. Eventually, the larva sinks to the seafloor and metamorphoses into the adult form.
While metamorphosis is the usual state in molluscs, the cephalopods differ in exhibiting direct development: the hatchling is a 'miniaturized' form of
the adult.[36] The development of molluscs is of particular interest in the field of ocean acidification as environmental stress is recognized to affect the
settlement, metamorphosis, and survival of larvae.[37]

Ecology

Feeding

Most molluscs are herbivorous, grazing on algae or filter feeders. For those grazing, two feeding strategies are predominant. Some feed on
microscopic, filamentous algae, often using their radula as a 'rake' to comb up filaments from the sea floor. Others feed on macroscopic 'plants' such
as kelp, rasping the plant surface with its radula. To employ this strategy, the plant has to be large enough for the mollusc to 'sit' on, so smaller
macroscopic plants are not as often eaten as their larger counterparts.[38] Filter feeders are molluscs that feed by straining suspended matter and
food particles from water, typically by passing the water over their gills. Most bivalves are filter feeders, which can be measured through clearance
rates. Research has demonstrated that environmental stress can affect the feeding of bivalves by altering the energy budget of organisms.[37]

Cephalopods are primarily predatory, and the radula takes a secondary role to the jaws and tentacles in food acquisition. The monoplacophoran
Neopilina uses its radula in the usual fashion, but its diet includes protists such as the xenophyophore Stannophyllum.[39] Sacoglossan sea-slugs suck
the sap from algae, using their one-row radula to pierce the cell walls,[40] whereas dorid nudibranchs and some Vetigastropoda feed on
sponges[41][42] and others feed on hydroids.[43] (An extensive list of molluscs with unusual feeding habits is available in the appendix of GRAHAM, A.
(1955). "Molluscan diets" (http://mollus.oxfordjournals.org/content/31/3-4/144.short). Journal of Molluscan Studies. 31 (3–4): 144..)

Classification
Opinions vary about the number of classes of molluscs; for example, the table below shows seven living classes,[19] and two extinct ones. Although
they are unlikely to form a clade, some older works combine the Caudofoveata and Solenogasters into one class, the Aplacophora.[27][15]: 291–292  Two
of the commonly recognized "classes" are known only from fossils.[17]

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Described living
Class Major organisms Distribution
species[19]
all snails and slugs including abalone, limpets, conch,
Gastropoda [15]: 300  nudibranchs, sea hares, sea butterflies
70,000 marine, freshwater, land

[15]: 367  clams, oysters, scallops, geoducks, mussels, rudists† 20,000 marine, freshwater
Bivalvia
Polyplacophora
[15]: 292–298  chitons 1,000 rocky tidal zone and seabed

Cephalopoda squid, octopuses, cuttlefish, nautiluses, Spirula,

[15]: 343  900 marine
belemnites†, ammonites†

Scaphopoda
[15]: 403–407  tusk shells 500 marine 6–7,000 metres (20–22,966 ft)

† Cricoconarida extinct

Aplacophora
[15]: 291–292  worm-like molluscs 320 seabed 200–3,000 metres (660–9,840 ft)

Monoplacophora seabed 1,800–7,000 metres (5,900–23,000 ft); one species 200

[15]: 298–300  ancient lineage of molluscs with cap-like shells 31
metres (660 ft)

Rostroconchia†[44] fossils; probable ancestors of bivalves extinct marine

Helcionelloida†[45] fossils; snail-like molluscs such as Latouchella extinct marine

Classification into higher taxa for these groups has been and remains problematic. A phylogenetic study suggests the Polyplacophora form a clade
with a monophyletic Aplacophora.[46] Additionally, it suggests a sister taxon relationship exists between the Bivalvia and the Gastropoda. Tentaculita
may also be in Mollusca (see Tentaculites).

Evolution

Fossil record

Good evidence exists for the appearance of gastropods (e.g., Aldanella),

cephalopods (e.g., Plectronoceras, Nectocaris) and bivalves (Pojetaia,
Fordilla) towards the middle of the Cambrian period, c.
500  million years ago, though arguably each of these may belong only to
the stem lineage of their respective classes.[47] However, the evolutionary
The use of love darts by the land snail
history both of the emergence of molluscs from the ancestral group
Monachoides vicinus is a form of sexual
Lophotrochozoa, and of their diversification into the well-known living and
selection
fossil forms, is still vigorously debated.
The enigmatic Kimberella quadrata
(fossil pictured) from the Ediacaran Debate occurs about whether some Ediacaran and Early Cambrian fossils
has been described as being really are molluscs. Kimberella, from about 555 million years ago, has been described by some paleontologists as
"mollusc-like" because of its "mollusc-like",[48][49] but others are unwilling to go further than "probable bilaterian",[50][51] if that.[52]
features which are shared with
modern day molluscs. There is an even sharper debate about whether Wiwaxia, from about 505 million years ago, was a mollusc, and
much of this centers on whether its feeding apparatus was a type of radula or more similar to that of some
polychaete worms.[50][53] Nicholas Butterfield, who opposes the idea that Wiwaxia was a mollusc, has written
that earlier microfossils from 515 to 510 million years ago are fragments of a genuinely mollusc-like radula.[54] This appears to contradict the concept
that the ancestral molluscan radula was mineralized.[55]

However, the Helcionellids, which first appear over 540 million years ago in Early Cambrian rocks from
Siberia and China,[56][57] are thought to be early molluscs with rather snail-like shells. Shelled molluscs
therefore predate the earliest trilobites.[45] Although most helcionellid fossils are only a few millimeters
long, specimens a few centimeters long have also been found, most with more limpet-like shapes. The
tiny specimens have been suggested to be juveniles and the larger ones adults.[58]

Some analyses of helcionellids concluded these were the earliest gastropods.[59] However, other scientists The tiny
Spirally coiled
are not convinced these Early Cambrian fossils show clear signs of the torsion that identifies modern Helcionellid fossil
shells appear in
gastropods twists the internal organs so the anus lies above the head.[15]: 300–343 [60][61] Yochelcionella is
many
thought to be an [15]: 300–34
gastropods.
Volborthella, some fossils of which predate 530 million years ago, was long thought to be a cephalopod, early mollusc[45]
but discoveries of more detailed fossils showed its shell was not secreted, but built from grains of the
mineral silicon dioxide (silica), and it was not divided into a series of compartments by septa as those of
fossil shelled cephalopods and the living Nautilus are. Volborthella's classification is uncertain.[62] The Middle    = Septa
Cambrian fossil Nectocaris is often interpreted as a cephalopod with 2 arms and no shell, but the Late Cambrian
fossil Plectronoceras is now thought to be the earliest undisputed cephalopod fossil, as its shell had septa and a    = Siphuncle
siphuncle, a strand of tissue that Nautilus uses to remove water from compartments it has vacated as it grows, and
which is also visible in fossil ammonite shells. However, Plectronoceras and other early cephalopods crept along
the seafloor instead of swimming, as their shells contained a "ballast" of stony deposits on what is thought to be
the underside, and had stripes and blotches on what is thought to be the upper surface.[63] All cephalopods with
external shells except the nautiloids became extinct by the end of the Cretaceous period 65 million years ago.[64]
However, the shell-less Coleoidea (squid, octopus, cuttlefish) are abundant today.[65] Septa and siphuncle in nautiloid
shell
The Early Cambrian fossils Fordilla and Pojetaia are regarded as bivalves.[66][67][68][69] "Modern-looking"
bivalves appeared in the Ordovician period, 488  to  443 million years ago.[70] One bivalve group, the rudists,
became major reef-builders in the Cretaceous, but became extinct in the Cretaceous–Paleogene extinction event.[71] Even so, bivalves remain
abundant and diverse.

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The Hyolitha are a class of extinct animals with a shell and operculum that may be molluscs. Authors who suggest they deserve their own phylum do
not comment on the position of this phylum in the tree of life.[72]

Phylogeny

The phylogeny (evolutionary "family tree") of molluscs is a controversial subject. In ⁠

addition to the debates about whether Kimberella and any of the "halwaxiids" were ⁠Brachiopods

molluscs or closely related to molluscs,[49][50][53][54] debates arise about the

relationships between the classes of living molluscs.[51] In fact, some groups ⁠Bivalves
traditionally classified as molluscs may have to be redefined as distinct but related.[75]

Molluscs are generally regarded members of the Lophotrochozoa,[73] a group defined by ⁠Monoplacophorans
⁠("limpet-like", "living fossils")
having trochophore larvae and, in the case of living Lophophorata, a feeding structure
called a lophophore. The other members of the Lophotrochozoa are the annelid worms
and seven marine phyla.[76] The diagram on the right summarizes a phylogeny ⁠
⁠Scaphopods (tusk shells)
presented in 2007 without the annelid worms. ⁠⁠
⁠⁠
Because the relationships between the members of the family tree are uncertain, it is ⁠Gastropods
difficult to identify the features inherited from the last common ancestor of all ⁠(snails, slugs, limpets, sea hares)
⁠
molluscs.[77] For example, it is uncertain whether the ancestral mollusc was metameric ⁠Mollusca
⁠
(composed of repeating units)—if it was, that would suggest an origin from an annelid- ⁠
⁠
like worm.[78] Scientists disagree about this: Giribet and colleagues concluded, in 2006, Cephalopods
⁠
the repetition of gills and of the foot's retractor muscles were later developments,[12] ⁠(nautiloids,ammonites, octopus,
while in 2007, Sigwart concluded the ancestral mollusc was metameric, and it had a foot ⁠Lophotrochozoa squid, etc.)
⁠
used for creeping and a "shell" that was mineralized.[51] In one particular branch of the ⁠
family tree, the shell of conchiferans is thought to have evolved from the spicules (small ⁠
⁠Aplacophorans
spines) of aplacophorans; but this is difficult to reconcile with the embryological origins ⁠(spicule-covered, worm-like)
of spicules.[77] ⁠
⁠

⁠
The molluscan shell appears to have originated from a mucus coating, which eventually ⁠Polyplacophorans (chitons)
stiffened into a cuticle. This would have been impermeable and thus forced the
development of more sophisticated respiratory apparatus in the form of gills.[45] ⁠
Eventually, the cuticle would have become mineralized,[45] using the same genetic ⁠Wiwaxia

machinery (engrailed) as most other bilaterian skeletons.[78] The first mollusc shell
⁠† Halwaxiids
⁠
almost certainly was reinforced with the mineral aragonite.[28] ⁠ ⁠
⁠Halkieria
⁠

The evolutionary relationships within the molluscs are also debated, and the diagrams
⁠
below show two widely supported reconstructions: ⁠ † Orthrozanclus

⁠
⁠ ⁠ ⁠ † Odontogriphus
⁠Solenogastres ⁠ Solenogastres
⁠
⁠
A possible "family tree" of molluscs (2007).[73][74] Does not include
⁠Aculifera ⁠ ⁠ annelid worms as the analysis concentrated on fossilizable "hard"
⁠Caudofoveata ⁠ Caudofoveata
⁠ features.[73]

⁠ ⁠
⁠Polyplacophorans ⁠Polyplacophorans

⁠ ⁠
⁠Monoplacophorans ⁠Monoplacophorans

⁠Molluscs ⁠Molluscs ⁠
⁠ ⁠ ⁠
⁠ ⁠
⁠Bivalves ⁠Bivalves
⁠Testaria
⁠
⁠ ⁠ ⁠Scaphopods
⁠Scaphopods
⁠Conchifera
⁠ ⁠ ⁠
⁠ ⁠
⁠ ⁠
⁠ ⁠
⁠Gastropods ⁠Gastropods

⁠ ⁠
⁠Cephalopods ⁠Cephalopods

The "Aculifera" hypothesis[73] The "Testaria" hypothesis[73]

Morphological analyses tend to recover a conchiferan clade that receives less support from molecular analyses,[79] although these results also lead to
unexpected paraphylies, for instance scattering the bivalves throughout all other mollusc groups.[80]

However, an analysis in 2009 using both morphological and molecular phylogenetics comparisons concluded the molluscs are not monophyletic; in
particular, Scaphopoda and Bivalvia are both separate, monophyletic lineages unrelated to the remaining molluscan classes; the traditional phylum
Mollusca is polyphyletic, and it can only be made monophyletic if scaphopods and bivalves are excluded.[75] A 2010 analysis recovered the traditional
conchiferan and aculiferan groups, and showed molluscs were monophyletic, demonstrating that available data for solenogastres was
contaminated.[81] Current molecular data are insufficient to constrain the molluscan phylogeny, and since the methods used to determine the
confidence in clades are prone to overestimation, it is risky to place too much emphasis even on the areas of which different studies agree.[82] Rather
than eliminating unlikely relationships, the latest studies add new permutations of internal molluscan relationships, even bringing the conchiferan
hypothesis into question.[83]

Human interaction

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For millennia, molluscs have been a source of food for humans, as well as important luxury goods, notably pearls, mother of pearl, Tyrian purple dye,
sea silk, and chemical compounds. Their shells have also been used as a form of currency in some preindustrial societies. A number of species of
molluscs can bite or sting humans, and some have become agricultural pests.

Uses by humans

Molluscs, especially bivalves such as clams and mussels, have been an important food source since at least the advent of anatomically modern
humans, and this has often resulted in overfishing.[84] Other commonly eaten molluscs include octopuses and squids, whelks, oysters, and
scallops.[85] In 2005, China accounted for 80% of the global mollusc catch, netting almost 11,000,000 tonnes (11,000,000 long tons; 12,000,000
short tons). Within Europe, France remained the industry leader.[86] Some countries regulate importation and handling of molluscs and other
seafood, mainly to minimize the poison risk from toxins that can sometimes accumulate in the animals.[87]

Most molluscs with shells can produce pearls, but only the pearls of bivalves and some gastropods, whose shells
are lined with nacre, are valuable.[15]: 300–343, 367–403  The best natural pearls are produced by marine pearl
oysters, Pinctada margaritifera and Pinctada mertensi, which live in the tropical and subtropical waters of the
Pacific Ocean. Natural pearls form when a small foreign object gets stuck between the mantle and shell.

The two methods of culturing pearls insert either "seeds" or beads into oysters. The "seed" method uses grains of
ground shell from freshwater mussels, and overharvesting for this purpose has endangered several freshwater
mussel species in the southeastern United States.[15]: 367–403  The pearl industry is so important in some areas,
significant sums of money are spent on monitoring the health of farmed molluscs.[88]
Saltwater pearl oyster farm in
Seram, Indonesia Other luxury and high-status products were made from molluscs. Tyrian purple,
made from the ink glands of murex shells, "fetched its weight in silver" in the fourth
century BC, according to Theopompus.[89] The discovery of large numbers of Murex
shells on Crete suggests the Minoans may have pioneered the extraction of "imperial purple" during the Middle Minoan
period in the 20th–18th centuries BC, centuries before the Tyrians.[90][91] Sea silk is a fine, rare, and valuable fabric
produced from the long silky threads (byssus) secreted by several bivalve molluscs, particularly Pinna nobilis, to attach
themselves to the sea bed.[92] Procopius, writing on the Persian wars circa 550 CE, "stated that the five hereditary
satraps (governors) of Armenia who received their insignia from the Roman Emperor were given chlamys (or cloaks)
made from lana pinna. Apparently, only the ruling classes were allowed to wear these chlamys."[93]

Mollusc shells, including those of cowries, were used as a kind of money (shell money) in several preindustrial societies.
However, these "currencies" generally differed in important ways from the standardized government-backed and -
controlled money familiar to industrial societies. Some shell "currencies" were not used for commercial transactions, but
Byzantine Emperor
mainly as social status displays at important occasions, such as weddings.[94] When used for commercial transactions, Justinian I clad in Tyrian
they functioned as commodity money, as a tradable commodity whose value differed from place to place, often as a result purple and wearing
of difficulties in transport, and which was vulnerable to incurable inflation if more efficient transport or "goldrush" numerous pearls
behavior appeared.[95]

Bioindicators

Bivalve molluscs are used as bioindicators to monitor the health of aquatic environments in both fresh water and the marine environments. Their
population status or structure, physiology, behaviour or the level of contamination with elements or compounds can indicate the state of
contamination status of the ecosystem. They are particularly useful since they are sessile so that they are representative of the environment where
they are sampled or placed.[96] Potamopyrgus antipodarum is used by some water treatment plants to test for estrogen-mimicking pollutants from
industrial agriculture.

Harmful to humans

Stings and bites

Some molluscs sting or bite, but deaths from mollusc venoms total less than 10% of those from jellyfish stings.[98]

All octopuses are venomous,[99] but only a few species pose a significant threat to humans. Blue-ringed octopuses in the
genus Hapalochlaena, which live around Australia and New Guinea, bite humans only if severely provoked,[97] but their
venom kills 25% of human victims. Another tropical species, Octopus apollyon, causes severe inflammation that can last
for over a month even if treated correctly,[100] and the bite of Octopus rubescens can cause necrosis that lasts longer than
one month if untreated, and headaches and weakness persisting for up to a week even if treated.[101]

All species of cone snails are venomous and can sting painfully when handled,
although many species are too small to pose much of a risk to humans, and only a few
fatalities have been reliably reported. Their venom is a complex mixture of toxins,
some fast-acting and others slower but deadlier.[102][98][103] The effects of individual The blue-ringed octopus's
cone-shell toxins on victims' nervous systems are so precise as to be useful tools for rings are a warning signal;
this octopus is alarmed, and
research in neurology, and the small size of their molecules makes it easy to
its bite can kill.[97]
synthesize them.[102][104]

Disease vectors
Live cone snails can be dangerous
to shell collectors, but are useful to
neurology researchers.[102]

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Schistosomiasis (also known as bilharzia, bilharziosis or snail fever), a disease caused by the fluke worm
Schistosoma, is "second only to malaria as the most devastating parasitic disease in tropical countries. An
estimated 200 million people in 74 countries are infected with the disease – 100 million in Africa alone."[105] The
parasite has 13 known species, two of which infect humans. The parasite itself is not a mollusc, but all the species
have freshwater snails as intermediate hosts.[106]

Pests
Skin vesicles created by the
Some species of molluscs, particularly certain snails and slugs, can be serious crop pests,[107] and when
penetration of Schistosoma.
introduced into new environments, can unbalance local ecosystems. One such pest, the giant African snail
(Source: CDC)
Achatina fulica, has been introduced to many parts of Asia, as well as to many islands in the Indian Ocean and
Pacific Ocean. In the 1990s, this species reached the West Indies. Attempts to control it by introducing the
predatory snail Euglandina rosea proved disastrous, as the predator ignored Achatina fulica and went on to
extirpate several native snail species instead.[108]

See also
Land snail
Sea slug
Sea snail

Explanatory notes
a. The formerly dominant U.K. spelling mollusk is still used in the U.S.—see the reasons given by Gary Rosenberg (1996).[2] For the spelling
mollusc, see the reasons given in: Brusca & Brusca. Invertebrates (2nd ed.)..

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Further reading
Sturm, C.; Pearce, T.A. & Valdes, A. The Mollusks: A Guide to Their Study, Collection, and Preservation. Universal Publishers. 2006. 454 pages.
ISBN 1581129300
Trigo, J.E.; Díaz Agras, G.J.; García-Álvarez, O.L.; Guerra, A.; Moreira, J.; Pérez, J.; Rolán, E.; Troncoso, J.S. & Urgorri, V. (2018). Troncoso,
J.S., Trigo, J.E. & Rolán, E., ed. Guía de los Moluscos Marinos de Galicia. Vigo: Servicio de Publicacións da Universidade de Vigo. 836 pages.
ISBN 978-84-8158-787-6

External links
"Mollusca" (https://www.eol.org/pages/2195) at the Encyclopedia of Life
Researchers complete mollusk evolutionary tree; 26 October 2011 (https://phys.org/news/2011-10-mollusk-evolutionary-tree.html)
Hardy's Internet Guide to Marine Gastropods (https://conchology.be/?t=261)
Rotterdam Natural History Museum (http://www.nmr-pics.nl/) Shell Image Gallery
Mussel Watch Programme (https://web.archive.org/web/20150907012135/http://ccma.nos.noaa.gov/about/coast/nsandt/musselwatch.aspx)
Online biomonitoring of bivalve activity, 24/7: MolluSCAN eye (http://molluscan-eye.epoc.u-bordeaux1.fr/index.php?rubrique=accueil&lang=en)
Archived (https://web.archive.org/web/20161113173444/http://molluscan-eye.epoc.u-bordeaux1.fr/index.php?rubrique=accueil&lang=en) 2016-
11-13 at the Wayback Machine

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